Evolutionary diversity peaks at mid-elevations along an Amazon-to-Andes elevation gradient

Elevation gradients present enigmatic diversity patterns, with trends often dependent on the dimension of diversity considered. However, focus is often on patterns of taxonomic diversity and interactions between diversity gradients and evolutionary factors, such as lineage age, are poorly understood. We combine forest census data with a genus level phylogeny representing tree ferns, gymnosperms, angiosperms, and an evolutionary depth of 382 million years, to investigate taxonomic and evolutionary diversity patterns across a long tropical montane forest elevation gradient on the Amazonian flank of the Peruvian Andes. We find that evolutionary diversity peaks at mid-elevations and contrasts with taxonomic richness, which is invariant from low to mid-elevation, but then decreases with elevation. We suggest that this trend interacts with variation in the evolutionary ages of lineages across elevation, with contrasting distribution trends between younger and older lineages. For example, while 53% of young lineages (originated by 10 million years ago) occur only below ∼1,750 m asl, just 13% of old lineages (originated by 110 million years ago) are restricted to below ∼1,750 m asl. Overall our results support an Environmental Crossroads hypothesis, whereby a mid-gradient mingling of distinct floras creates an evolutionary diversity in mid-elevation Andean forests that rivals that of the Amazonian lowlands

An annotated checklist of trees and relatives in tropical montane forests from southeast Peru: the importance of continue collecting

Los Andes están considerados como los puntos calientes más diversos de los trópicos, dentro de estos se encuentra el Parque Nacional del Manu, cuyas complejas condiciones climáticas y fisiográficas albergan una mega-diversidad y endemismo. En el presente trabajo se presenta una lista anotada de especies arbóreas y afines a lo largo de un gradiente de elevación desde los bosques submontanos a 800 m hasta la línea de bosque a 3625 m en la Reserva de Biosfera del Manu. En base a una red de 21 parcelas permanentes de una hectárea y exploraciones botánicas se sistematiza la información florística por rangos de elevación, distribución geográfica y endemismo. Estos resultados preliminares se traducen en 1108 especies de las cuales el 39.7% son morfoespecies, el 43% de las especies determinadas son registros nuevos para la región del Cusco, 15 especies son nuevos registros para la flora peruana, 40 especies son endémicas para Perú y 30 son potenciales especies nuevas para la ciencia. Adicionalmente, se resalta la expansión del rango altitudinal para el 45.2% de las especies determinadas (302 especies). Estos resultados son una muestra de la alta diversidad arbórea y afines en estos ecosistemas montañosos registrados en tan solo ~20 km de distancia geográfica, además muestra lo escasamente colectados y poco estudiados que se encuentran. Mas colecciones botánicas son necesarias - estos estudios básicos de florística son imperativos para un mejor entendimiento de la distribución de especies y la función del ecosistemas, además ayudará a responder una de las grandes preguntas en la ecología global moderna, ¿Cómo responderán los bosques tropicales al cambio climático global?The tropical Andes and adjacent Amazon are Earth’s highest biodiversity hotspot. Manu National Park in southeastern Peru encompasses an entire watershed, ranging from Andean highlands to Amazonian lowlands, and is a megadiverse landscape on the Andes to Amazon transition. Here we present an annotated checklist of trees and related species is along an elevation gradient in the Manu Biosphere Reserve that runs from sub-montane forests at 800 m elevation up to the tree line at 3625 m. Based on a network of 21 1-hectare permanent tree plots and botanical explorations, the floristic information is systematized by elevation ranges, geographical distribution and endemism. These preliminary results show 1108 species. Of these, 43% are new records for the region of Cusco, 15 species are new records for the Peruvian flora, 40 species are endemics for Peru, and 30 are potential new species for science. Another 39.7% are identified to genus or family level and remain morphospecies. Additionally, we show altitudinal range expansion for 45.2% of identified species (302 species). These results were found in a transect of plots spanning only 20 km of geographic distance, and are a sample of the high tree diversity in these mountainous ecosystems. The data show how poorly collected and understudied these ecosystems are. Basic floristic studies and collections are imperative for a better understanding of species distribution and function of ecosystems, and the basic biodiversity of the tropical Andes. They will also help to answer a major, unresolved question in modern global ecology of how tropical forests will respond to global climate change

Phylogenetic diversity of Amazonian tree communities

This is the peer reviewed version of the following article: Honorio Coronado, E. N., Dexter, K. G., Pennington, R. T., Chave, J., Lewis, S. L., Alexiades, M. N., Alvarez, E., Alves de Oliveira, A., Amaral, I. L., Araujo-Murakami, A., Arets, E. J. M. M., Aymard, G. A., Baraloto, C., Bonal, D., Brienen, R., Cerón, C., Cornejo Valverde, F., Di Fiore, A., Farfan-Rios, W., Feldpausch, T. R., Higuchi, N., Huamantupa-Chuquimaco, I., Laurance, S. G., Laurance, W. F., López-Gonzalez, G., Marimon, B. S., Marimon-Junior, B. H., Monteagudo Mendoza, A., Neill, D., Palacios Cuenca, W., Peñuela Mora, M. C., Pitman, N. C. A., Prieto, A., Quesada, C. A., Ramirez Angulo, H., Rudas, A., Ruschel, A. R., Salinas Revilla, N., Salomão, R. P., Segalin de Andrade, A., Silman, M. R., Spironello, W., ter Steege, H., Terborgh, J., Toledo, M., Valenzuela Gamarra, L., Vieira, I. C. G., Vilanova Torre, E., Vos, V., Phillips, O. L. (2015), Phylogenetic diversity of Amazonian tree communities. Diversity and Distributions, 21: 1295–1307. doi: 10.1111/ddi.12357, which has been published in final form at 10.1111/ddi.12357Aim: To examine variation in the phylogenetic diversity (PD) of tree communities across geographical and environmental gradients in Amazonia. Location: Two hundred and eighty-three c. 1 ha forest inventory plots from across Amazonia. Methods: We evaluated PD as the total phylogenetic branch length across species in each plot (PDss), the mean pairwise phylogenetic distance between species (MPD), the mean nearest taxon distance (MNTD) and their equivalents standardized for species richness (ses.PDss, ses.MPD, ses.MNTD). We compared PD of tree communities growing (1) on substrates of varying geological age; and (2) in environments with varying ecophysiological barriers to growth and survival. Results: PDss is strongly positively correlated with species richness (SR), whereas MNTD has a negative correlation. Communities on geologically young- and intermediate-aged substrates (western and central Amazonia respectively) have the highest SR, and therefore the highest PDss and the lowest MNTD. We find that the youngest and oldest substrates (the latter on the Brazilian and Guiana Shields) have the highest ses.PDss and ses.MNTD. MPD and ses.MPD are strongly correlated with how evenly taxa are distributed among the three principal angiosperm clades and are both highest in western Amazonia. Meanwhile, seasonally dry tropical forest (SDTF) and forests on white sands have low PD, as evaluated by any metric. Main conclusions: High ses.PDss and ses.MNTD reflect greater lineage diversity in communities. We suggest that high ses.PDss and ses.MNTD in western Amazonia results from its favourable, easy-to-colonize environment, whereas high values in the Brazilian and Guianan Shields may be due to accumulation of lineages over a longer period of time. White-sand forests and SDTF are dominated by close relatives from fewer lineages, perhaps reflecting ecophysiological barriers that are difficult to surmount evolutionarily. Because MPD and ses.MPD do not reflect lineage diversity per se, we suggest that PDss, ses.PDss and ses.MNTD may be the most useful diversity metrics for setting large-scale conservation priorities.FINCyT - PhD studentshipSchool of Geography of the University of LeedsRoyal Botanic Garden EdinburghNatural Environment Research Council (NERC)Gordon and Betty Moore FoundationEuropean Union's Seventh Framework ProgrammeERCCNPq/PELDNSF - Fellowshi

Bryophyte stable isotope composition, diversity and biomass define tropical montane cloud forest extent.

Liverworts and mosses are a major component of the epiphyte flora of tropical montane forest ecosystems. Canopy access was used to analyse the distribution and vertical stratification of bryophyte epiphytes within tree crowns at nine forest sites across a 3400 m elevational gradient in Peru, from the Amazonian basin to the high Andes. The stable isotope compositions of bryophyte organic material (13C/12C and 18O/16O) are associated with surface water diffusive limitations and, along with C/N content, provide a generic index for the extent of cloud immersion. From lowland to cloud forest δ13C increased from -33‰ to -27‰, while δ18O increased from 16.3‰ to 18.0‰. Epiphytic bryophyte and associated canopy soil biomass in the cloud immersion zone was estimated at up to 45 t dry mass ha-1, and overall water holding capacity was equivalent to a 20 mm precipitation event. The study emphasizes the importance of diverse bryophyte communities in sequestering carbon in threatened habitats, with stable isotope analysis allowing future elevational shifts in the cloud base associated with changes in climate to be tracked.NERC PhD Studentshi

Phylogenetic diversity of Amazonian tree communities

Aim: To examine variation in the phylogenetic diversity (PD) of tree communities across geographical and environmental gradients in Amazonia. Location: Two hundred and eighty-three c. 1 ha forest inventory plots from across Amazonia. Methods: We evaluated PD as the total phylogenetic branch length across species in each plot (PDss), the mean pairwise phylogenetic distance between species (MPD), the mean nearest taxon distance (MNTD) and their equivalents standardized for species richness (ses.PDss, ses.MPD, ses.MNTD). We compared PD of tree communities growing (1) on substrates of varying geological age; and (2) in environments with varying ecophysiological barriers to growth and survival. Results: PDss is strongly positively correlated with species richness (SR), whereas MNTD has a negative correlation. Communities on geologically young- and intermediate-aged substrates (western and central Amazonia respectively) have the highest SR, and therefore the highest PDss and the lowest MNTD. We find that the youngest and oldest substrates (the latter on the Brazilian and Guiana Shields) have the highest ses.PDss and ses.MNTD. MPD and ses.MPD are strongly correlated with how evenly taxa are distributed among the three principal angiosperm clades and are both highest in western Amazonia. Meanwhile, seasonally dry tropical forest (SDTF) and forests on white sands have low PD, as evaluated by any metric. Main conclusions: High ses.PDss and ses.MNTD reflect greater lineage diversity in communities. We suggest that high ses.PDss and ses.MNTD in western Amazonia results from its favourable, easy-to-colonize environment, whereas high values in the Brazilian and Guianan Shields may be due to accumulation of lineages over a longer period of time. White-sand forests and SDTF are dominated by close relatives from fewer lineages, perhaps reflecting ecophysiological barriers that are difficult to surmount evolutionarily. Because MPD and ses.MPD do not reflect lineage diversity per se, we suggest that PDss, ses.PDss and ses.MNTD may be the most useful diversity metrics for setting large-scale conservation priorities

Estimating the global conservation status of more than 15,000 Amazonian tree species

Estimates of extinction risk for Amazonian plant and animal species are rare and not often incorporated into land-use policy and conservation planning. We overlay spatial distribution models with historical and projected deforestation to show that at least 36% and up to 57% of all Amazonian tree species are likely to qualify as globally threatened under International Union for Conservation of Nature (IUCN) Red List criteria. If confirmed, these results would increase the number of threatened plant species on Earth by 22%. We show that the trends observed in Amazonia apply to trees throughout the tropics, and we predict thatmost of the world’s >40,000 tropical tree species now qualify as globally threatened. A gap analysis suggests that existing Amazonian protected areas and indigenous territories will protect viable populations of most threatened species if these areas suffer no further degradation, highlighting the key roles that protected areas, indigenous peoples, and improved governance can play in preventing large-scale extinctions in the tropics in this century

Wood density variation across an Andes‐to‐Amazon elevational gradient

Understanding how functional traits are related to species diversity and ecosystem properties is a central goal of ecology. Wood density is a trait that integrates many aspects of plant form and function and is highly variable among species. Previous studies of wood density across elevational gradients have been based on limited sampling and have reported declines with increasing elevation, though even this simple pattern remains unknown, much less its underlying functional and evolutionary relationships. Here, we use one of the longest and most speciose elevational gradients in the world, extending from the Andean tree line to the Amazon basin, to test the extent to which elevation, species composition, phylogenetic affinity and forest structure determine variation in wood density. Using field‐collected wood samples and global databases, we assigned wood density to 1231 species and 31,330 stems across 41 (47.5 ha) mature forest plots arrayed across a 3500‐m vertical gradient. Our results show that mean wood density, either weighted by abundance, basal area or species, was highly variable but tended to decline from low to middle elevations and increase again from mid‐elevations to the tree line. As a result of this non‐linearity, forests at the Andean tree line had higher wood density than their lowland Amazon counterparts. We observed an abrupt transition in wood density at the lower limit of persistent cloud formation (cloud base), where the lowest wood density values were found. The decline of wood density is attributed to a significant shift in life forms, with an abundance of tree ferns at middle elevations and a higher probability of landslides and disturbances favouring a suite of traits associated with low wood density, such as softer wood and higher elasticity. Species turnover explained most of the among‐species variation across the gradient, with elevation having no consistent effect on within‐species variation in wood density. Together, both gradual compositional changes and sharp local changes in the importance of non‐dicot life forms, such as arborescent ferns and palms, define patterns of forest‐level carbon density, with wood density per se controlling ecosystem properties, such as carbon flux, across the Andes‐to‐Amazon elevational gradient

The variation of productivity and its allocation along a tropical elevation gradient: a whole carbon budget perspective

Why do forest productivity and biomass decline with elevation? To address this question, research to date generally has focused on correlative approaches describing changes in woody growth and biomass with elevation. We present a novel, mechanistic approach to this question by quantifying the autotrophic carbon budget in 16 forest plots along a 3300 m elevation transect in Peru. Low growth rates at high elevations appear primarily driven by low gross primary productivity (GPP), with little shift in either carbon use efficiency (CUE) or allocation of net primary productivity (NPP) between wood, fine roots and canopy. The lack of trend in CUE implies that the proportion of photosynthate allocated to autotrophic respiration is not sensitive to temperature. Rather than a gradual linear decline in productivity, there is some limited but nonconclusive evidence of a sharp transition in NPP between submontane and montane forests, which may be caused by cloud immersion effects within the cloud forest zone. Leaf-level photosynthetic parameters do not decline with elevation, implying that nutrient limitation does not restrict photosynthesis at high elevations. Our data demonstrate the potential of whole carbon budget perspectives to provide a deeper understanding of controls on ecosystem functioning and carbon cycling

Modern pollen rain predicts shifts in plant trait composition but not plant diversity along the Andes–Amazon elevational gradient

This is the final version. Available on open access from Wiley via the DOI in this record. Aims: Terrestrial ecosystems are changing in biodiversity, species composition and functional trait composition. To understand the underlying causes of these changes and predict the long-term resilience of the ecosystem to withstand future disturbances, we can evaluate changes in diversity and composition from fossil pollen records. Although diversity can be well estimated from pollen in temperate ecosystems, this is less clear for the hyperdiverse tropics. Moreover, it remains unknown whether functional composition of plant assemblages can be accurately predicted from pollen assemblage composition. Here, we evaluate how community-weighted mean (CWM) traits and diversity indices change along elevation. Location: Amazon–Andes elevation gradient in Peru. Methods: We used 82 modern pollen samples and 59 vegetation plots along the elevation gradient, and calculated CWM traits and diversity indices for each pollen sample and vegetation plot. We also quantified the degree to which taxa are over- or underrepresented by their pollen, by dividing the relative pollen abundance by the relative basal area abundance in the nearby vegetation survey plots (i.e. the R-rel values). Results: We found that CWM wood density increased, and CWM adult height and leaf area decreased with elevation. This change was well predicted by pollen assemblages, indicating that CWM trait–environment relationships based on pollen abundance data provide meaningful results. Diversity (richness, Shannon and Simpson) decreased with elevation for vegetation plots, but these trends could not be observed from pollen assemblages. Conclusions: Our results demonstrate that more research is needed to develop methods that lead to accurate diversity estimates from pollen data in these tropical ecosystems, but that CWM traits can be calculated from pollen data to assess spatial shifts in functional composition. This opens opportunities to calculate CWM traits from fossil pollen data sets in the tropics, with broad implications for improving our understanding and predictions of forest dynamics, functioning and resilience through time.Nederlandse Organisatie voor Wetenschappelijk Onderzoe

Plant trait and vegetation data along a 1314 m elevation gradient with fire history in Puna grasslands, Perú

Alpine grassland vegetation supports globally important biodiversity and ecosystems that are increasingly threatened by climate warming and other environmental changes. Trait-based approaches can support understanding of vegetation responses to global change drivers and consequences for ecosystem functioning. In six sites along a 1314 m elevational gradient in Puna grasslands in the Peruvian Andes, we collected datasets on vascular plant composition, plant functional traits, biomass, ecosystem fluxes, and climate data over three years. The data were collected in the wet and dry season and from plots with different fire histories. We selected traits associated with plant resource use, growth, and life history strategies (leaf area, leaf dry/wet mass, leaf thickness, specific leaf area, leaf dry matter content, leaf C, N, P content, C and N isotopes). The trait dataset contains 3,665 plant records from 145 taxa, 54,036 trait measurements (increasing the trait data coverage of the regional flora by 420%) covering 14 traits and 121 plant taxa (ca. 40% of which have no previous publicly available trait data) across 33 families